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\3 Biodiversity 

Transactions of the Zoological Society of London. 

[London] Published for the Zoological Society of London by Academic Press 

V. 4 1862: 

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[ 31 ] 

III. On the Anatomy of ike Indian Rhinoceros (Rh. unicornis, L.) 

By Professor Owen, F.R.S^ F.Z*S. Sfc. 

Read Feb. 12, 1S50. 

Part I. 

Introduction. External characters. Position of Viscera. 

XHE very rare opportunity of investigating the internal structure of the Rhinoceros, 
which the death of the fine male specimen of the Indian species, Rhinoceros unicornis $ L-, 
at the Menagerie of the Zoological Society, has afforded, enables me to submit to the 
Society the following details of its anatomy. 

I may premise, as a requisite point of comparison with the dimensions and weight of 
some of the viscera, that the animal, which was full-grown and had lived in the mena- 
gerie fifteen years, measured thirteen feet and a half from the end of the muzzle to the 
root of the tail, and thirteen feet in its greatest circumference ; its total weight was 
upwards of two tons 1 . 

The animal had begun to show a loss of appetite in July 1849, when it was supposed 
to be under the influence of the rut : the more decided symptoms of ailment first mani- 
fested themselves about a week before its death, when it was observed to make occa- 
sional efforts, as if to vomit, followed by the escape of a bloody and frothy mucus and 
fluid from the mouth and also from the nose. It died on the evening of the 19th of 
November 1849. Subjoined are the symptoms noted in the Head- Keeper's Minute- 
book 5 * 

After the removal of the integuments and some dissection of the muscles, the abdo- 
minal and thoracic viscera were exposed by the detachment of all the ribs of the left 
side ; when it was found that the seventh rib had been fractured at the bend near the 
vertebral end : a kind of false joint had been formed between the broken portions. One 

Mr, Miller, the Superintendent, has transmitted to me a record which shows that the Rhinoceros, when 
received at the Gardens, 20th September 1834, weighed I J ton: there was no means of weighing the entire 
animal after Its death : bat an approximation was made by weighing separately the limbs, the trunk, detached 
masses of flesh, the hide, &c. t which allowed the total weight to be estimated at about 5000 lbs. avoirdupois. 
J 849, November ]2th. Rhinoceros vomited slimy mucus. 

■s *t 




with blood. 








ditto, and from the nostril-. 











1 9th. 





VOL, IV. — PART 11. 




of these had wounded the left lung, and the inflammation, which had caused extensive 
adhesion of the back part of the lung to the pleura costalis, had also extended into the 
pulmonary substance, and along the bronchial tubes into the trachea. The surface of 
the part of the left lung near the wound was extensively emphysematous ; and the in- 
flamed bronchial tubes were loaded with bloody frothy serum and mucus. The supposed 
attempts at vomiting were doubtless efforts to disembarrass the windpipe of the successive 
accumulations of this fluid ; and the death of the animal is to be ascribed to the injury 
and disease of the left lung consequent on the fracture. 

The other morbid appearances were of minor moment: a portion of the right lung 
was the seat of Hydatids, of the genus Echinococcus. The parent cysts were of various 
sizes from the diameter of two inches to that of half an inch ; two or three being suc- 
cessively included in one another. The uncinated vermicules floating freely in the fluid 
of the parent cysts were xiHioth of an inch in diameter, and in countless numbers. 
They will be more particularly described by Mr. Qnekett in the Appendix to this paper ; 
in which also will be given the particulars of the morbid state of the gastric follicles in 
the digesting portion of the stomach , as observed by the microscope : to this state may 
be attributed the failure of appetite which first drew attention to the declining health of 
this rare and valuable quadruped. 

The calcareous matter which was discovered in the gastric follicles would probably 
have laid the basis of a gastric or intestinal calculus, if the animal had lived ; but the 
apparently healthily digested condition of a considerable proportion of the contents of 
the stomach showed that the state of the secreting apparatus could only be remotely 
connected with the last fatal symptoms. The liver was less firm than usual ; but this 
might be due to the rapidity with which the large pachyderms pass into a state of 
chemical decomposition after death. There is a striking difference in that respect 
in different mammalia ; the Ruminants resist the decomposing forces longer than the 
Pachyderms, as I have experienced in dissecting the Giraffe and the Aurochs j and 
the resistance is more remarkable in some other orders. On dissecting the two-toed 
Sloth in moderately warm weather, I was surprised at the length of time in which it 
kept sweet. Martin, in his ■ History of the British Colonies,' observes of the Sea Cow 
(Manatus americanus), "Its flesh is white and delicate, resembling veal in appearance 
and taste, and it will keep good several weeks, even in the hot climate of which it is a 
native, when other meat will not resist putrefaction for as many days," The Elephant 
and the Tapir which I have dissected at the Society's Gardens rapidly passed, like the 
hlliinoeeroSj ipto an offensive state of decomposition. 

The bodies of the 3th, 6th and 7th dorsal vertebra were anchylosed together along 
their nnder part, from which an exostosis of apparently old growth projected into the 
base of the mediastinum, forming there an obtnse rounded tumour of about two inches 
vertical thickness and twelve inches in circumference. The neural arches and spines of 
these vertebras showed no fracture or disease, and as there had not been any symptom 



of paralysis, further damage to the skeleton for the purpose of examining the spinal 
marrow at that part was not deemed expedient. 

Our able and active Secretary has reminded me, that at the time when the large male 
Elephant was exhibited along with the Rhinoceros in a contiguous paddock, the latter 
used to submit to be poked on the back by the Elephant, who could lift his head over 
the palings and press down with his tusks upon the thick hide of the Rhinoceros ; and 
that the Rhinoceros has been observed to have been thus forced down until his belly 
touched the ground. Now although this procedure did not actually fracture the spines 
or neural arches so pressed upon, it most probably strained the ligaments beneath the 
bodies of the same vertebrae, and produced the ossific inflammation which led to the 
anchylosis and tumour discovered on dissection, One cannot, however, attribute to 
this old injury the immediate cause of the animal's rapidly fatal malady. It may have 
led to the fracture of the rib articulating to the anchylosed parts, through the sup- 
pression of that degree of elastic yielding, which the interspace of the vertebrae in their 
ordinarily moveable state would have afforded. The animal, in lying down, usually 
fell heavily on its side, and the rib had probably become fractured on one of these 
occasions by £ centre-coup.* 

The external form and characters of the present specimen of Indian Rhinoceros 
agreed with the full and often-repeated descriptions which have been already published ; 
especially with that excellent one by Dauhenton in Buffon's * Histoire Naturelle/ 4to, 
torn. xL p. 198, and with the description, illustrated by two fine and accurate figures, 
by F. Cuvier in the 'Histoire Naturelle des Mamtnif£res, s foL, fasc, xiiL 1820, p. 2. 
The only point which appears to have escaped these and other good observers, is the 
orifice behind each carpus and tarsus, which forms the termination of the duct of a 
pretty large subdermal glandular pouch. 

In the male Rhinoceros unicornis, in a state of nature, the horn, when the animal has 
attained the length of nine feet eight inches, is, according to Mr. Hodgson, the learned 
and accomplished Resident at Nepal, five inches in height. In the Society's specimen, 
the horn, owing to the habit which the animal had acquired of rubbing and beating it 
against the woodwork of its den, had never been permitted to grow beyond eight inches 
in height ; but its base measured nine inches in transverse breadth, eleven inches in 
anteroposterior extent, and twenty-six inches in circumference. It was distant 



From the inner canthus of the eye 

From the end of the upper lip . 

From the occipital ridge . - * - 

In a female Rhinoceros of the same species which died in a travelling menagerie in 
January 1838 1 , and was purchased by the Royal College of Surgeons, the horn was a mere 

1 This animal was found dead in its den after a night during which the thermometer had fallen IO p below 

G 2 



callous protuberance, scarcely an inch in height and only four inches across the base, 
although the animal was nine feet in length and the same in its greatest circumference. 
This difference indicates that the epidermal production in question varies in its pro- 
portions in the male and female, making a sexual distinction analogous to that which 
may be observed in the antlers of the Reindeer, the false antlers of the Giraffe, and in 
some other Ruminants in which horns are present in both sexes 1 . 

Since, as Daubenton well demonstrated, and as has been amply confirmed and eluci- 
dated by later observers, the horn of the Rhinoceros is an unvascular production, an 
agglutination of fibres like bristles, unsupported by any osseous core, the question early 
suggested itself to me, how the relative position, of £he epidermal conglomerate to the 
eye and the end of the muzzle was preserved during the progressive growth of the head, 
and I have carefully watched the progress of the horn in the male animal here described 
from its first reception into the Society's Menagerie. During the whole of the period 
of the animal's growth, the back part of the horn was that which alone exhibited natural 
decay ; the fibres there being ragged and broken, while the new fibres were added at the 
sides and chiefly in front. Thus the horn kept pace, as to its relative position, with 
the progressive elongation of the jaws during the acquisition of the permanent teeth, by 
a process analogous to that by which the adductor muscle of the oyster maintains the 
same relative position to the hinge and outlet of the shell during the whole period of 
the shell's growth. This partial or local decay and renovation became less conspicuous 
after the Rhinoceros had attained its full size, and in the long and large horns of aged 
individuals the whole circumference presents the same smooth and polished surface: 
whence it may be concluded that when the skull, and especially the upper jaw of the 
Rhinoceros have attained their full size, the horn receives additional matter along the 
whole extent of the base, and increases more rapidly in length than in the immature 

The glandular orifices at the back part of each foot to which I have alluded, are 
situated about three inches above the callous sole in the fore-feet and about two and a 
half inches above the sole in the hind-feet : they are concealed from cursory observation 
in the middle of the transverse fold that runs parallel to the interspace between the 
carpus and metacarpus, and between the tarsus and metatarsus. The orifice is analo- 
gous to that which opens on the tore part of the foot between the digits in the Sheep 
and some other Ruminants. The gland itself (PL IX, fig. 1) in the Rhinoceros is of a 
compressed ovate figure, measuring one and a halt inch in length and one inch in 
breadth. The thickness of the glandular parietes (/ft. fig. 2) varies from two to three 
lines. These parietes consist of a compact congeries of follicles, surrounded externally 

the freezing point : it had previously exhibited no signs of disease, and had been carried about and exhibited 
upwards of a year. 

1 In the female Rhinoceros Simus the anterior horn is longer and more slender than In the male: in other 
two-horned species I am not at present aware of any sexual distinction in these weapons. 



a muscular and tendinous coat : the diameter of the excretory orifice {lb. fig. 3) 
is about eight lines when fully expanded. 

The anus is dilatable to a great extent, corresponding to the large masses in which 
the feces are discharged ; and the Rhinoceros presents, in the size of this aperture, the 
opposite extreme to the Giraffe and the Nilghau. 

A deep groove formed by two thick and prominent parallel folds extends from below 
the anus along the median line of the perinaeum, gradually widening, to the buck part 
of the base of the preputial or pendulous part of the penis. In the ordinary retracted 
state of this organ, the distance from the anus to the preputial orifice is two feet ten 
inches : the length of the thick, wrinkled, tegumentary prepuce (PL IX. fig, 4), within 
which the long glans is commonly retracted and concealed, is about nine inches ; 
its basal circumference measures one foot six inches ; on each side of this part there is 
a hemispheroid warty prominence (lb. a). When the animal stales, the glans is pro- 
truded downwards with a curve backwards {Ik fig, 5), and the urine is ejected between 
the hind-lee;s in a succession of jerks. I never saw it issue in a continuous stream. 
Under sexual excitement the penis is much further protruded and drawn forwards 
in a straight line (lb. fig. 6). The singular form of the glans will be subsequently 


In the female Indian Rhinoceros, the vagina and the preputium clitoridis open on the 
external surface by two separate, narrow, vertically elongated orifices, each being, in the 
individual of nine feet long— the subject of the remarks on this sex in the present 
Memoir, about two inches in length : the vulva {lb. fig. 7, vu) is immediately above 
the clitoris {lb. fig. 8, e), i.e. nearer the anus. The extent of the perineum between the 
anus and vulva was four and a half inches. 

The nipples {lb. fig. 8, m, m) were two in number and inguinal ; they were situated 
fourteen inches in advance of the vulva, and two and a half inches apart from one 
another. They were subcompressed, obtusely rounded at the extremity, and about two 
inches in length : about a dozen lactiferous ducts opened upon the somewhat flattened 

summit of each nipple. 

The integument on the middle line of the abdomen, along which it was divided in the 
operation of skinning, presented a general thickness of three-fourths of an inch : where 
it was cut on the inner side of the extremities, it was about one-fourth of an inch in 
thickness. It was connected to the abdominal parietes by a loose cellular tissue, and 
by a closer subcutaneous tissue to most of the other parts of the body; but the parts to 
which the stiff and ponderous hide most firmly adhered were the spinous processes of 
the posterior lumbar and sacral vertebrae, and the anterior extremities of the iliac bones, 
at which places the corium was blended with the periosteum, and was remarkably thin. 
The hide adhered over the jugal bones to a kind of moveable fibro-cartilage ■ and its 
attachment along the median line of the fore part of the head was so firm as to require, 
especially beneath the horn, the use of a chisel in order to separate it from the skull 



But, besides its attachment to subcutaneous cellular tissue, fasciae, elastic tissue, 
iibro-cartilages and periosteum, the hide is connected with parts which are destined for 
its motions and adjustment upon the body. So far from the panniculus carnosus being 
absent, it is developed in certain parts to an extraordinary thickness; and it became 
obvious, on contemplating these muscles, that one use of the permanent folds in the 
hide of this thick-skinned species of Rhinoceros, is to afford, like the processes of bones, 
a firmer insertion to the aponeuroses of the cutaneous muscles than a plane surface of 
integument could possibly have done. A sheet of panniculus carnosus situated on each 
side of the thoracic or scapular region sends its fascia into the interstice of the fold in 
front of the anterior extremities, the skin being bent upon itself, as it were, to grasp 
this fascia. Similar portions of panniculus carnosus send their aponeuroses into the 
posterior folds of the skin. But the most remarkable portions of the cutaneous muscu- 
lar system are two, which arise, broad and thick, one on each side of the anterior part 
of the abdomen from the superficial fascia covering that part, and, passing backwards, 
terminate in aponeurotic sheets which are inserted into the fasciae covering the patelhe 
and knee-joint. As the patellae are higher than the abdomen, in the erect position of 
the animal, the preceding muscles would seem to be developed chiefly to afford additional 
support to the bulky abdomen, the weight of w r hich is thus in part transferred imme- 
diately to the hinder extremities ; and these the muscles in question must also tend to 
draw forwards during progressive motion* 

The dense but highly elastic e fascia superficial^ \ spread over the peripheral surface 
of the abdominal muscles upon their pubic and hypoehondrial regions, increases in 
thickness as it passes over the abdominal rings, and invests the spermatic chord with a 
thick sheath, which becomes thinner where it expands upon the ■ tunica vaginalis testis. 5 
Each testis was situated out of the abdomen, but pretty close to the external abdominal 
ring, without, however, causing any protuberance in the thick integument : and there 
is no scrotum or outward indication of the essential glands of the male organs. 

In the female the superficial fascia covering the external abdominal rings descended 
upon and surrounded the mammary glands ; which occupy a corresponding position to 
that of the testes in the male 1 * On the internal or central surface of the mammary 
glands was situated a plexus of large veins : the arteries supplying them were a branch 
from the superficial femoral and branches of an artery answering to the cremasteric 
artery in the male, which passed, with the ■ ligamentum teres uteri * through each abdo- 
minal ring. 

The soie of each foot was occupied by a thick cushion of elastic tissue, not adipose 
chiefly, as in Man, but of a whiter, gelatinous and ligamentous texture, resembling 
the morbid tissue called £ albuminous carcinoma* 1 The difference between the thick 
epidermal layer covering the sole, and that sheathing the fore part of each of the three 

1 This correspondence is accompanied by a similarity in the development and functions of the c remaster 
muscle in the two sexes of the Marsupial quadrupeds, 



toes, is more marked than in the Horse : the hoofs proper, or homologues of the nails, 
are firmly attached to the periosteum of the ungual phalanges by fine vertical lamina 
interlocking with corresponding vascular laminae of the thickened periosteum. 

Before entering on the subject of the visceral anatomy of the Rhinoceros unicornis, I 
may premise that some general details on this subject will be found in a paper by 
Dr. James Parsons in the Philosophical Transactions for 1743, on the occasion of the 
death of the Rhinoceros sent by the Chief of the Hon. E. L Company from Patna to 
London in the year 1739 i I possess an impression of a scarce print of the animal pub- 
lished in London in that year. 

A second Rhinoceros of the same species, which was exhibited and died in London 
in 1800, was dissected by Honoratns Leigh Thomas, Esq,, who has given an account of 
his observations on that occasion in a paper printed in the Philosophical Transactions 
for 180 L Mr, William Bell had previously contributed to the Philosophical Transactions 
for 1793, some interesting remarks on the anatomy of the Sumatran two-horned Rhino- 
ceros, then for the first time described. 

In the one-horned Indian specimens dissected by me, the peritoneal membrane was 
thick and much stronger than in the human subject : the cellular tissue connecting the 
external surface of this serous membrane to the adjacent structures is condensed into 
an aponeurotic firmness where it is attached to the serous coat, the free surface of which 
presents an opake, whitish appearance. In the female Rhinoceros I exposed the abdo- 
minal viscera by laying open the cavity along the middle line of the ventral surface, and 
turning aside the flaps of its yielding softparietes. Not the least trace of epiploon was 
observable when the cavity of the abdomen was thus exposed ■ but the viscera which 
presented themselves were in immediate contact with the sustaining parietes, A single 
but enormous fold of the colon, not less than two feet in breadth, formed more than one 
half of the exposed surface of the abdominal viscera ; it passed obliquely across the 
middle of the cavity, from the right hypochondriac to the left hypogastric or iliac region ; 
immediately below this was a smaller fold of colon 1 running parallel with the preceding ; 
below this was a second fold ; and, occupying the right iliac region, a part of the smooth 
parietes of the caecum appeared : a portion of the liver and the stomach were obscurelv 
visible in the epigastric and hypochondriac regions, and below these were seen a few 
coils of the small intestine. 

The colon was not displaced without considerable difficulty, owing to the weight of 
its contents, and the strength of the duplicatures of the peritoneum attaching it to the 
spine and contiguous parts, Behind and above the great oblique folds of colon lay a 
short, thin and corrugated epiploon* devoid of fat; and behind and below them were 
several coils of the small intestines : the spleen and kidneys were also brought into 

1 It is to these enormous folds of this colon that the great size of the abdomen is due, and not to the caecum, 
which is not proportionally so large as in the Horse. 



view j together with the large caecum, appearing like a second stomach , occupying the 
right iliac and lumbar regions. 

In the male Rhinoceros the thoracic and abdominal viscera were exposed by the suc- 
cessive detachment of the ribs of the left side, together with the soft walls of the same 
side of the thorax and abdomen. The diaphragm separating these two cavities extended 
from about the seventeenth dorsal vertebra obliquely downwards and forwards* curving, 
as it approached the ventral parietes, more rapidly towards them ; its diameter following 
this course being four feet six inches. The length of the abdominal cavity was seven 
feet ; its depth or antero-posterior diameter three feet six inches. The length of the 
thoracic cavity near the spine was three feet six inches ; its depth at the most promi- 
nent part of the convex diaphragm was two feet ; its size, contrasted in this view with 
that of the enormous abdomen } seemed disproportionately small. 

The viscera of the abdomen which presented themselves, enumerated from the dia- 
phragm backwardSs were the free curved border and part of the upper convex surface 
of the left lobe of the liver, partly overlapping the stomachy of w r hich about two-thirds 
of the greater or cardiac portion were visible. The lower free border of the spleen ex- 
tended from below all the visible part of the great curvature of the stomach ; and the 
thin, fatless, shrivelled epiploon was continued from beneath the spleen upon the upper 
part of the base of the great fold of the colon above mentioned. This enormous fold 
slipped forwards as soon as the supporting walls of the abdomen were removed, and 
exposed the large coils of the left descending portion of the colon continued from it, 
and below and ventrad of these were exposed some of the coils of the small intestine. 
A part of the left kidney protruding at the angle between the cardiac end of the stomach 
and the commencement of the descending colon, was covered by a duplicature of peri- 
toneum extending from its ventral surface to the contiguous end of the spleen* 

The dorsal border of the left lobe of the liver was attached by a similar duplicature , 
forming a strong * ligamentum triangulare ' to the contiguous part of the diaphragm. 
The length of the great fold of the colon taken in a straight line as it lay first exposed 
was six feet six inches : some idea of its capacity may be formed from the fact that the 
portion of the fold next the caecum could easily contain a man, with ample room for him 
to turn about in it But the dimensions of the alimentary canal and its several parts 
will be subsequently given. 



Part II. 

Digestive Organs. Abdominal Viscera, 

The Mouth. — The substance of both the lower and upper lip was composed of cellular 
and subliganientous tissue permeated in all directions by muscular fibres, and resembling 
in section the ( corpus cavernosum penis 1 in the Horse : the skin covering this substance 
is very thin and vascular in the upper lip. These muscular fibres, which are homolo- 
gous with the decussating fibres in the proboscis of the Elephant, presented the striated 
characteristic of the voluntary muscular fibre under the microscope. 

The seventh pair of nerves , which was lost principally in the muscles and the above- 
described contractile tissue of the upper lip, was of large size. 

In the male Rhinoceros the tongue measured two feet three inches from the epiglottis 
to the tip, and seven and a half inches across its broad anterior part : the depth or 
thickness of the tongue is four inches, at its root. In the female Rhinoceros the tongue 
measured nineteen inches in length from the epiglottis to the tip. This organ is broad 
and fiat, slightly expanded at its anterior extremity, and becoming narrower and deeper 
as it extends backwards : there is a small protuberance on the upper surface opposite the 
posterior grinders, divided by a longitudinal depression : the large fossulate papillae of the 
dorsum are principally collected in a group of ten to twelve on each of these risings : 
the epithelium is disposed on the anterior part of the tongue in a number of very fine 
close-set pointed papillae, resembling short hairs: behind the papillae the epithelium is 
condensed into a thick callous stratum, which gradually becomes thinner where it covers 
the posterior glandular part of the tongue. There are no retroverted cuticular processes, 
as in the Ruminants. There is a lytta beneath the anterior flattened part of the tongue, 

A reticulate structure at the sides of the soft palate, having muciparous follicles in 
the interspaces of the meshes, and many subcompressed conical processes of various 
lengths, represents the tonsils (PL X. t t t t t) : the arches of the palate, or * isthmus 
faucium,* form on each side a thin sharp fold, which descends obliquely along the sides 
of the pharynx and terminates insensibly near the sides of the glottis. The soft palate 
consists of a stratum of muciparous follicles one-third of an inch thick, placed ver- 
tically between two layers of mucous membrane ; their blind extremities being in contact 
with the whitish dense membrane lining the nasal or air-passage, their orifices termi- 
nating on the soft red and vascular membrane at the roof of the mouth. The constrictors 
of the pharynx formed at the anterior margin of that canal a thick rounded edge. 

The pointed apex of the triangular epiglottis (16. e) curves forward above the base 
of the tongue, to which the epiglottis is attached by a pair of strong c glosso-epi- 
glottidei ' muscles. 

The alimentary canal — The oesophagus extends pretty straight from the pharynx to 
the stomach, with an uniform diameter, in its passive or contracted state, of three 
inches : its total length was five feet. It extends about six inches into the abdomen 




the cardi 

fice about 

foot five 

after piercing the diaphragm, and terminates s 
inches from the left extremity of the stomach. This organ [PI. XL figs. 1 & 2) pre- 
sented the ordinary form of the simple stomach; it was moderately distended with 
food; with a large obtuse cardiac end, expanding to the cardiac orifice (fig. 2, c), 
opposite to which it presented the greatest circumference ; thence contracting to near 
the pylorus (ib. p), on the cardiac side of which the stomach presented its smallest 
circumference ■ and then expanding into a blind end, of a hemispheric form, beyond 
the pylorus. The length of the stomach in a straight line was four feet ; its diameter 
from the cardia to the opposite part of the great curvature was one foot ten inches. 
The small curvature between the cardia and pylorus was one foot nine inches. There 
was a glistening aponeurotic sheet (ib. a) upon the anterior and posterior surfaces of 
the contracted pyloric end of the stomach. 

A sheet of white thick epithelium spreads from the cardia over the inner surface of 
the cardiac portion of the stomach, about one foot four inches along the lesser cur- 
vature, and along the greater curvature to the extent shown in figure 2, e. This epi- 
thelial layer is one line thick, smooth, or with very fine rugse on its inner surface, and 
terminating by a well-defined border, near which it is perforated by numerous orifices 
of mucous follicles (lb, fig. 4), The rest of the inner surface of the stomach pre- 
sents the usual vascular structure, with the more minute orifices of the secerning fol- 
licles of the gastric juice. There is no crescentic fold or valve at the cardia, as in the 
Horse : nor is there any valvular protuberance on the gastric side of the pylorus, as in 
the Cow and most other Ruminants : the thickened rim of the pylorus was slightly 
produced into the duodenum. 

In the female Rhinoceros the stomach presented the same simple elongated form as 
in the male, corresponding with the description of its external form given by Cuvier 
(after Vicq, d'Azyr?) 1 , Its total length in a straight line was thirty-two inches, and 
the distance from the cardia to the left extremity was fourteen inches. It was distended 
with a mass of coarsely divided hay mixed with oats. The whole of the cardiac extre- 
mity, excepting at one small spot, was lined with a smooth compact layer of thickened 
epithelium, like that in the male : it extended along the upper or smaller curvature of 
the stomach half-way between the cardia and the pylorus ■ its greatest extent from the 
left end of the stomach being twenty-two inches. The boundary-line between this and 
the glandular or mucous coat of the stomach was even, but as abrupt and well-marked 
as in the Horse. The epidermis was very easily detached^ and in some places had sepa- 
rated spontaneously, as does the thick epithelial lining of a gizzard soon after death, and 
it is probable that such spontaneous separation of the cuticle in the Rhinoceros dissected 
by Mr. Thomas may have induced the belief that it was wanting in that animal •. The 

1 Lemons d'Anat. Corop* ifL (1805) p. 392. 

3 "The stomach upon its inside was in every part covered by a secreting surface; whereas in the Horse it 
h partly cuticukr." — Pirihs* Tmns, 1S01, p, 14-7* 



Indian species agrees, however, in the twofold nature of the lining membrane of the 
stomach, with the Sumatran two-horned Rhinoceros described by Mr. William Bell 1 . 
About the middle of the cuticuiar surface of the stomach of the female there was a small 
irregular patch of glandular membrane: this was proved to be an original formation, 
and not an appearance due to a partial separation of the cuticle, by detaching the 
surrounding cuticuiar lining and comparing the patch in question with the denuded 
surface. It is probably, however, but an individual variety, as it was not repeated in 
the male. The surface of the digestive membrane covering the pyloric moiety of the 
stomach was even, not broken by mgse, and it presented the same peculiar smooth, 
almost polished, appearance which characterizes the peculiar glandular membrane lining 
the second cavity of the stomach of the Porpoise. 

The cardia did not present the semi-spiral valve observable in the Horse. The 
globular pyloric extremity is suddenly bent upon the rest of the stomach, so as to ap- 
pear partly separated from it by the entering fold, A thick circular lip projects from 
the pylorus into the duodenum. The outer layer of the muscular tunic, a, is one- 
fourth the thickness of the inner layer, 6, and becomes thinner over the pyloric end of 
the stomach. The nervous or vasculo^cellular tunic, c, begins to increase in thickness 
near the termination of the thick epithelium, d, in relation to the increased vascular 
action required by the functions of the glandular layer, e : the relative thickness of this 
laver is shown in the section, figure 3, c'. 

The contents of the duodenum were of a greenish black colour and almost fluid con- 
sistency : only very few small portions of the vegetable substances appeared in the tract 
of the small intestines, but the caecum and colon were tensely distended with a magma 
of substances like those in the stomach, but of somewhat softer consistence, as if in a 

further stage of digestion* 

50 feet. 

8 inches, 

6 inches. 

7 inches. 


65 feet, 
10 inches. 

8 inches. 

9 inches. 

The length of the small intestines was • - . 
The circumference of the duodenum .... 

The circumference of jejunum ...... 

The circumference of ileum 

The lining membrane of the duodenum, at the beginning of that gut, was puckered 
up into small irregular rugBe ; the flattened triangular processes, as described and figured 
by Mr. Thomas, began to make their appearance about six inches from the pylorus 
(PI. XII. fig, 1) ; in the jejunum three or four of the processes are often supported on 
a common base ( lb. fig. 2) ; as they approach the ileum they begin to lose breadth, and 
gain in length, until they assume the appearance, near the end of the ileum, of vermi- 
form processes, like tags of worsted, from two-thirds of an inch to an inch in length 
{lb, fig. 3). Peyer's glands appeared scattered here and there- a very conspicuous 
reticular patch was situated close to the end of the ileum. 

1 Philos. Trans. 1793* Cuvier does not describe the inner surface of stomach. 

H 2 



The small intestines have nearly the same disposition as in the Horse ; they are sus- 
pended by a short mesentery, in which the anastomosing arteries form only one series of 
arches. The mucous membrane of the ileum projects in the form of a circular fold within 
the caecum ; but it seems inefficient as a valve for preventing regurgitation of at least 
fluid matters from the large intestines. The length of the caecum (Pi. XIII. cm) from this 
orifice to its blind extremity in the male Rhinoceros was three feet, and its greatest cir- 
cumference was four and a half feet. In the female Rhinoceros the length of the caecum 
was two feet ; its circumference two feet six inches ; these proportions to the colon and 
the rest of the intestinal canal being rather less than in the Horse, The anterior sur- 
face of the ci-ecum is traversed longitudinally by a fibrous band, four inches broad, upon 
which it is slightly sacculated \ a second band appears, nearer the colon. Its lining 
membrane was puckered up into innumerable irregular small transverse rugge, which 
appear, however, to be but temporary foldings of the mucous membrane, and are easily 
obliterated when this is stretched. The colon for the first four feet of its extent was 
puckered up upon three longitudinal bands into sacculi, each about five inches long : it was 
suddenly bent upon itself at this part, forming the long and large fold {lb. co\ co J ), the 
two parts of the fold being very closely connected to each other ; it there became dilated 
into the very wide portion which formed the most prominent object on laying open the 
abdomen ; the beginning of this dilated portion is also closely adherent by its posterior 
surface to the opposite surface of the beginning of the csecum. The circumference of 
this dilated part of the colon (which if permanent, and not due to accidental accumu- 
lation of alimentary matter, might be regarded as representing a second ceecutn or 
reservoir,) is five feet : beyond this fold the colon becomes gradually narrower, its 
smallest circumference being twenty inches, where it passes into the rectum , which 
forms several short convolutions before its termination. 



19 feet. 

25 feet, 

3 feet. 

5 feet. 

The entire length of the colon was . . - 
The entire length of the rectum . 

The total length of the intestinal canal, including the cascum, was in the female 
seventy-three feet; in the male ninety-six feet, or eight times the length of the entire 

The circumference of the rectum was ten inches in the female, and sixteen inches in the 
male ; but it widens towards the anus. The masses in which the fseces are discharged 
from the immense receptacles formed by the large intestine, are greater than in the 
Elephant, and are softer and more amorphous. 

The longitudinal muscular fibres of the rectum were developed into such powerful 
fasciculi as to lead me to suspect some change of tissue ; but on examining the fibre 
microscopically, it presented the same absence of aggregation of the ultimate fibres into 
striated bundles, as in the higher tract of the intestines. The contrast between these 



fibres in the rectum and those of the external sphincter was well-marked, the latter pre- 
senting the striated character of true voluntary muscles. 

The herbivorous Mammalia differ from the carnivorous more in the character of their 
large intestines than of their small intestines. The less putrefactive nature of their food 
renders it susceptible of a longer retention in the body ; and the receptacular and sac- 
cular character of the large intestines seems especially designed to retard the course of 
the alimentarv substances. An observation made by the celebrated Surgeon Dupuytren, 
throws light upon the final purpose of this detention of the food of the Herbivora: he 
noticed in a patient who had an artificial anus near the end of the small intestines, 
that the vegetable parts of the food thence ejected were undigested. Dr. Beaumont 
also observed that the vegetable substances underwent much less change than the 
animal substances in the stomach of the man (Alexis) with the fistulous opening into 
the stomach. That organ in the artiodactyles (Peccari, Hippopotamus, and Ruminants) 
is rendered specially complex for overcoming the difficulty, and the cseeum and colon 
are comparatively small : but in the perissodactyles (Horse, Tapir, Rhinoceros) the 
more simple stomach is compensated by the increased capacity and complexity ot 
the lame intestines. The subdivided stomach in the Sloths is in some respects, as 
e. g. the glandular appendage, and vascular secerning surface of the paunch, more com- 
plex than that of Ruminants : and here accordingly we find the caecum absent and the 
colon undefined. These facts should be kept in mind by the Physiologist when he draws 
from Comparative Anatomy in support of inferences as to the special function of the 
cfficum in completing the digestion of vegetable food. The Dormouse and other hyber- 
nating Rodents are far from being the sole exceptions to the presence of a proportionally 
large aecum in the Herbivora : a large CEecum is rather the exception than the rule in the 
vegetable feeders. It is only found in those Herbivora, in which, through the necessity 
of a correlation with other circumstances than that of the nature of the food, the stomach 
retains the simple form and moderate size of that of the carnivorous or mixed feeding 
mammals. Comparative Anatomy significantly warns us against ascribing a special or 
exclusive importance to any particular dilatation of the alimentary canal. It plainly 
demonstrates that neither a complex stomach nor a large caecum are essential to the 
digestion of vegetable food ; but it teaches that a capacious and complex alimentary 
canal is essential for that purpose, at least in the Mammalia. Either a highly- deve- 
loped and concentrated glandular apparatus must be added to the stomach, as in the 
Dormouse, Wombat and Beaver ; or the stomach must be amplified, subdivided or sac- 
culated, as in the Ruminants and herbivorous Marsupials ; or both complexities must be 
combined, as in the Sloths, Dugongs and Manatees j or, if a simple condition of stomach 
is retained, it must be compensated by a large sacculated colon and oecum. 

Digestive glands.— The liver presented the dark colour noticed by Mr. Thomas in his 
dissection of the Rhinoceros, In the female specimen which I examined, its textore 
was as firm as in the Horse, and its weight was 21 lbs. avoirdupois. In the older and 



larger male its texture was softer and more grumous ; and it weighed 44 lbs. avoirdu- 
pois* With respect to its form, I did not find an agreement either with the statement 
of Mr. Thomas 1 or the description in the second edition of the ( Lemons d'Anatomie Com- 
paree, 3 iv. p. 464 (1836). In both specimens of Rhinoceros the liver was divided into 
fewer lobes than ordinary, taking the Mammalia generally, yet had a right lobe in addi- 
tion to the principal bifid lobe and the left lobe, the two latter only being assigned to it 
by Cuvier. The form of the gland is flattened, as in the hoofed animals generally , its 
greatest thickness was not more than six inches in the male. Its longest or transverse 
diameter measured in the female twenty-seven inches, and the length or antero-poste- 
rior diameter of the middle lobe seventeen inches. Three great hepatic veins join the 
inferior cava just below the diaphragm. The strong serous tunic of the liver was 
beautifully marked by arborescent vessels of a white colour. The ' ligamentum rotun- 
dum J and corresponding fold of peritoneum entered as usual into the notch dividing 
the middle lobe, which might be compared to the cystic lobe in the quadrupeds which 
possess a gall-bladder. This appendage, however, as in Mr, Thomas's dissection 2 , was 
wanting, as it is also in the other perissodactyle or odd-toed Pachyderms; e. j. the 
Hyrax, the Tapir, the Elephant, and the Horse. In these, as in the Rhinoceros, the 
absence of the gall-bladder seems to be dependent on the small size of the stomach as 
compared with the quantity of food taken, to the consequent frequency of feeding, and 
to the rapid and probably un intermitting transit of the gastric contents through the 
small intestines to the enormous caecal and colonic receptacles where digestion and 
animal izat ion are finally completed 3 . The great biliary duct is formed in the portal 
fissure by the union of six or seven branches from the lobes of the liver : its diameter is 
half an inch j it terminates in the duodenum six inches from the pylorus. 

The pancreas resembles that of the Horse and Tapir: its principal duct (PL XIV. 
fig. 1, h) enters the intestine close to the biliary duct (lb. a) } communicating therewith 
in the oblique course between the tunics r the duct of the smaller portion of the 
pancreas (lb, ft) terminates about two inches from the large and protuberant common 
opening of the preceding ducts, but at the same distance from the pylorus. 

The spleen is an elongated, subtrihedral, flattened body, lodged in the duplicatures of 
the short omentum. It weighed 5 lbs. in the male, and 3 lbs. in the female Rhinoceros: 
in the latter its length was two feet six inches ; its greatest breadth one foot ; its smallest 
breadth six inches: in the male it measured three feet six inches in lengthy one foot 
four inches in breadth: it resembles in structure that of the Horse. 

Kidneys,— -The weight of these two glands was about 8 lbs, in the female and 1 1 lbs. 
in the male Rhinoceros. In both they had the same situation in the abdomen as in the 
Horse, They were lobulated, and the extent of subdivision was intermediate between 

1 "It was divided into several lobes/' — Tom. cit* - See also Cuvier* Ice, cit iv* p. 519- 

1 See the excellent remarks by Mr, Youatt in Lis work on * The Horse,' Svo, 1831, p» 212. In Lhe Hog the 
esecum its comparatively small* 



that which respectively characterizes the kidneys of the Ox and Bear ; the average size 
of the component lobules being two inches 1 . In the female, the kidneys did not resemble 
each other in form. That on the left side was flattened and semi-ovate, ten inches long, 
six and a half inches broad, and two inches thick. The right kidney was subtrian- 
gular (PL XIV. fig. 2), presenting a flattened surface to the broadly expanded ribs on 
which it rested, and having on the opposite or anterior side two flattened surfaces 
meeting at an obtuse angle : this kidney was eight and a half inches long, seven and a 
half inches broad in the female. In the male the kidneys were more symmetrical : the 
right measured eleven inches in length and seven inches in breadth. The great vein, 
the artery, and the ureter had the usual relative position near the pelvis of the kidney ; 
the vein being anterior, and the ureter descending behind the artery : this duct 
presented a diameter of half an inch. 

The ureter (JS, fig. 3, u) having penetrated the substance of the gland for the extent of an 
inch t divides into two branches (J6. p, p) at right angles to the trunk : one branch ascends, 
the other descends, and both together form a long canal which may be called the ' pelvis* 
of the kidney. Into this canal the common trunks (/ft. t) of the radiating E tubuli uriniferi/ 
from the several lobes> open without forming any valvular protuberance or ' mammilla. 1 
There is the same facility, therefore, for injecting the * tubuli' as in the Horse or Tapir. 

A white injection of size and flake-white was thrown into the ureter, and forced 
into the tubuli uriniferi by pressing the injection onwards towards the kidney* and 
thus alternately emptying the ureter hy the finger and thumb, and filling the ureter 
from the syringe : the tubuli uriniferi were injected as far as the superficies of the gland j 
and the injection was continued until a few specks of extravasation appeared ; but not 
any portion of injection returned either by the artery or vein. 

In the right kidney the tubuli iiriniferi were tilled, with similar success, and afterwards 
the emulgent artery was injected with red size injection ; this returned by the vein, but 
did not penetrate any of the branches of the ureter. The tubuli uriniferi form loops at 
the periphery of the kidney, returning into the cortical substance* 

The ureters, which preserve a diameter of about half an inch through their whole 
course, penetrate the urinary bladder, in the male (at u 9 « } PL XVL}, a little w r ay above 
the fundus of the * vesiculse seminales,' where they are about six inches apart, but they 
converge in their oblique course through the thick muscular coat of the bladder. In the 
female they are more closely approximated at their terminations, which, in the young 
animal I dissected, were only half an inch apart, and about one line in diameter : their 
orifices were six inches from the commencement of the urethra. This short tube 
opened into the urogenital canal (PL XVII L fig. \ 7 u) five inches from the vulva. 

The Rhinoceros examined by Mr* Thomas had not attained its third year, which led that gentleman to 
conjecture that the tabulated structure might be lost as the animal advanced id life {loc* cit. p* 148) ; but the 
persistence of this struct a re to the ninth year, ia the female, and to the fifteenth year, in the male, animal dis- 
sected by me, proves that to be a permanent condition of the renal organ in the Rhiuuctros, as in smiie other 
Mammalia, which is a commuu foetal peculiarity in Man. 



Suprarenal glands.— These bodies, like the kidneys a differed from each other in form j 
they were elongated and nearly cylindrical The right had one extremity bent at a right 
an^te : its length in the female Rhinoceros was three and a half inches ; its breadth across 
the bent extremity two inches: the left was simply elongated, three and a half inches 
long, one and a half broad and one inch thick* In section they presented an external 
greyish-yellow fibrous cortex, from one-fourth to one-third of an inch thick, enclosing 
a fleshy-coloured substance, in the middle of which there was a semilunar portion of the 
grey fibrous matter : there was no trace of a centra! cavity. Both suprarenal bodies 
adhered closely to the contiguous large veins* 

The urinary bladder presented nothing remarkable except a very distinct pit or cica- 
trix, surrounded by a double concentric fold of membrane (PL XIV, fig* 4, d) r where 
the duct of the allantois originally communicated with the cavity. 

Part IIL 

Thoracic Viscera* 

The thoracic viscera presented much the same relative position as in the Horse; the 
lungs becoming narrow and elongated at the contracted anterior part of the thorax ; the 
distance between the pericardium and the diaphragm was relatively less than in the 


The heart weighed 28 lbs, avoirdupois. The length of the undistended ventricular 
part was one foot one inch ; the breadth of the ventricles was one foot three inches. 
The pericardium was of great strength* The heart presented the short, obtuse form 
which characterises it in the Elephant and Tapir, 

The superior precaval vena cava receives the right or common vena azygos close to 
its termination at the upper part of the right auricle : two inches above this it receives 
the right vertebral vein, which is about half an inch in diameter; two inches above 
this it is formed by the junction of the left subclavian with the right subclavian vein. 
At the concavity of the great vein formed by this junction, which concavity crosses the 
fore part of the aortic arch, the bronchial veins and some small pericardial veins enter the 
superior cava. The upper part of the superior cava receives the two large jugular veins 
close together, so that a proper 'vena innominata* can scarcely be said to be formed. 
The left vena azygos, which is formed by the union of a few superior intercostal veins 
of the same side, terminates in the left subclavian vein, which receives separately the 
left vertebral vein from the neck. The right or principal azygos receives the intercostal 
veins of both sides as far forwards as its entry into the precaval vein ; the Rhinoceros in 
this structure agreeing with the Horse, 

The coronary vein receives only a small pericardial vein, which descends along the 
back of the left auricle, before it terminates with the inferior cava, at the base of the 
right auricle. 



There was no trace of a valve at the orifices, either of the inferior cava or coronary 
vein ; the latter easily admitted the end of the fore-finger. In the right ventricle, the 
tricuspid valve presented the following attachments ; — its strong chordae tend mere were 
distributed to three obtuse and transversely oblong columnar earner one rising from the 
external or moveable wall, a second from the septum, and a third smaller one from the 
anterior interspace between the fixed and moveable wall : the tendons diverged from 
each column to the two contiguous moieties of the divisions of the tricuspid valve ; a 
provision which ensures the simultaneous action and the outstretching of these three 
membranous processes. There were besides two smaller columns placed opposite to 
each other, one on the free and the other on the fixed wall of the ventricle j they were 
connected together by a single strong tendon passing across the ventricular cavity from 
the apex of one to that of the other. 

The mitral and semilunar valves offered nothing unusual. 

The aorta, after giving off two coronaries, each of which freely admitted the fore- 
finger, ascended and divided at the summit of its arch into the descending aorta and 
a smaller trunk supplying the head and anterior extremities. The vessels immediately 
derived from the ascending division were the two internal thoracics 1 , the brachials, and 
the common trunk of the two carotids. 

Each lung was divided into a small upper and a large lower lobe ; the right lung gave 
off in addition a transversely elongated narrow azygos lobe. The superior lobe of each 
lung was characterized by numerous deep marginal notches, which gave it an appendicu- 
lated character. The lining membrane of the branches of the bronchia? presented very 
strongly marked longitudinal rugae; that of the trachea was similarly disposed. After 
reflecting the pleura from the surface of the lung, a thin extensible stratum of condensed 
cellular tissue continuous with the interlobular cellular tissue could alone be perceived. 
Between the pleuree and the parietes of the chest was much elastic tissue. 

The cartilaginous hoops of the trachea are stout and close-set ; they meet posteriorly, 
but their extremities do not coalesce; their number was 31. The diameter of the 
windpipe is two inches and a half, being not greater than that of the Lion* 

The larynx consists of the thyroid (PL XV. (A), cricoid {lb. c) f and arytenoid (lb, a) 
cartilages, of the epiglottis (16. e), and of a small sesamoid tibro-cartilage [lb. fig. 1, h) 
developed in the commissure of the * arytenoidei transversa and ' obliqui/ here blended 
together; but there is no trace of the cartilages of Santorin or Wrisberg. The wings 
of the thyroid cartilage meet at a slightly obtuse angle, contrary to their usual dis- 
position in the Hog tribe and Ruminants : there is no notch at the upper margin of 
the anterior median line ; but there is a considerable triangular vacancy below, filled 

1 The intercostal spaces above or anterior to the heart, are numerous in proportion to the narrowness of the 
chest, which obliges the heart to be placed nearer the diaphragm ; and the internal thoracics, which are of insig- 
nificant size in Man, are there largely developed in order to supply those intercostal spaces, which, from the 
position of the heart, cannot receive their arteries directly horn the trunk of the aorta. 

VOL* IV. — PART n, I 



up by dense elastic and aponeurotic membrane, to which yielding walls of the larynx 
some of the fibres of the thyreo-arytenoidei muscles adhere. The arytenoid cartilages 
are relatively of large size ; their base extends half-way across the aperture of the 
larynx, and from the anterior extremities of these produced bases, the upper 
(PL XV. fig. 1, u) and lower {Ih. v) 'chord* vocales' extend forwards to the thyroid 
cartilage and base of the epiglottis. Only the anterior half, therefore, of the ' rima 
glottidis' is bounded by vibratile vocalizing material, and the ordinary feeble bleat of 
the Rhinoceros (like that of a calf) is wbat might be expected to be produced by such 

a structure. 

On each side, between the upper and lower chordae vocales there is the opening of a 
large sacculus laryngis, which communicates anteriorly with a crescentic fossa under 
the base of the epiglottis. A fold of membrane (Pis. X* & XV. /) extends on each side 
from a small fibro~ca.rtilai>e (/}, at the inner or under side of the base of the epiglottis, 
downwards, inwards, and forwards to the anterior termination of the chordae vocales, u 
and v : these oblique folds form the inner or posterior walls of the anterior fossae of the 
sacculi laryngis. 

The anterior or superior labia (PL XV. fig. 2, m) of the glottis form two broad, thick, 
slightly everted folds of mucous membrane, 

Jn the mass of muscles (PI. XV. fig. 2, o, o) attached to and passing between the 
arytenoid cartilages, there are developed about twelve tendons which radiate to be 
inserted into the central sesamoid cartilage before mentioned. 

The epiglottis (Pis. X. & XV, e, e) is of a triangular figure, with the pointed apex 
curved forwards, and having strong glosso-epiglottidei muscles attached to it. 

The thyroid gland consisted of two elongate, subtriangular lobes extending from 
the sides of the larynx to the fourth tracheal ring; diminishing as they descend and 
united by a very thin and narrow strip continued between their inferior extremities^ 
obliquely across the front of the trachea. The structure of this body is more distinctly 
lobular than is usually seen ; a small compact yellow glandular body was attached to 
the thyroid at the point where the veins emerge. 

Part IV. 

Generative Organs. 

Male organs. — The cremaster is a very powerful muscle, and consists of coarse 
carneous fasciculi in two flattened masses, one crossing the other obliquely as they 
escape with the spermatic chord beneath the arch of the abdominal ring. A cluster of 
lymphatic glands with much tough elastic cellular tissue fill up the rest of the ring. 
The cremaster at this part measures one inch and a half in breadth and half an inch in 




The external inguinal position of the testis in close contact with the abdominal r 
has already been described. The tunica vaginalis communicated freely with the peri- 
toneal cavity. Each testis presented an oval figure, seven inches in length, four inches 
and a half in breadth, and four inches in thickness. It is surrounded by a strong and 
thick e tunica albuginea/ On making a section into the gland along the line of attach- 
ment of the epididymis, the * corpus Highmoriauum' was exposed, in the form of a 
moderately thick white band, continued from the end of the gland where the efferent 
vessels pass out to form the t caput epididymides, 1 along the whole longitudinal axis of 
the gland. From this almost ligamentous band or centre of the cellular framework of 
the gland, the septal layers diverge to all parts of the external tunic of the testicle, 
forming the compartments in which the lobes of aggregated £ tubuli seminiferi' are 
lodged. The branches of the spermatic artery, on penetrating the tunica albuginea, 
pass directly to the corpus Highmorianum, and their ramifications diverge thence, sup- 
ported by the radiating septa, and form a rich network upon the inner or vascular layer 
of the capsule of the testis. 

The vas deferens enters the inguinal canal surrounded by the vessels and especially 
by the plexiform veins of the spermatic chord, and oa entering the abdomen is received 
in a peritoneal, fold and is conducted to the side and then to the back part of the urinary 
bladder, passing between the bladder and the ureter: having got to the inner side of 
the termination of the ureter, the vasa deferentia (PL XVI + fig. I , vd t vd) descend straight , 
slightly converging, to the middle of the back part of the prostate : they penetrate that 
gland, together with the ducts of the vesiculpe seminales, lying to the inner side of these ; 
and, communicating with them, the common duct on each side finally terminates by a 
minute pore (PL XVII. fig. 4) upon the crucial verumontanum. The vasa deferentia are 
thickened to about thrice their ordinary diameter in the last three inches of their course ; 
but their canal or area is not proportionally dilated ; it is, on the contrary, rather 
contracted, by the thickness of the cellulo-glandular parietes to which the enlargement 
of the duct is due- 

The vesicular glands or ' vesicutae seminales* (PL XVI. fig* 1 , vs t m) present an elongate 
subcompressed pyriform shape, eight inches in length, and three inches and a half across 
the broadest part of the fundus. They have a lobulated exterior, and a structure very 
similar to that of the same bodies in Man, 

The prostate (lb. pr t pr) is much less compact than in Man and more resembles that 
of many Rodents, being composed of an aggregate of long slender csecai tubes with glan- 
dular walls, converging to the ducts of the vesicular and vasa deferentia, and opening by 
numerous minute apertures on the verumontanum (PI, XVIL fig, 4). The breadth of the 
prostate is six inches ■ its antero-posterior extent four inches : it does not quite surround 
the beginning of the urethra, but is closely applied to the back and sides of that canaL 

The muscular or membranous part of the urethra, m, extends about three inches from 
the prostate before it joins the bulbous and cavernous portions, close to which are 



situated two large subeompressed oval Cowperian glands (lb., c, c). Each of these 
measures three indies and a half by two inches and a half. The structure of the corpus 
cavernosum resembles that of the Horse. 

The great plexus of veins above the dorsum penis near its root, was enveloped in a 
mass of elastic tissue, like the ' dartos 1 of the human scrotum. 

The fleshy part of the ' levatores penis 1 (PL XVIL It) measures fourteen inches in length, 
five inches across their basal origin, and between one and two inches in thickness. 
Their oblique origin is extended over the space of one foot from the ento-pelvic part of 
the pubis down to the ischium. The tendinous part of the muscle commences where 
the pubic portion joins the ischial portion of the muscle at the inner and under border 
of the fleshy part ; it is half an inch thick at its commencement, but expands as it ex- 
tends along the muscle, the fleshy fasciculi of which are inserted into the tendon in an 
obliquely converging, or semUpenniform manner. As the tendon augments in breadth, 
it diminishes in thickness, converging towards its fellow, which it meets and joins two 
inches before the anterior termination of the fleshy portion, The two united flattened 
tendons beyond are gradually converted into a round chord of ligamentous substance an 
inch in diameter. This chord (lb. t) glides through a strong, slightly elastic aponeurotic 
sheath along the median groove of the dorsum penis ; it is connected with the inner 
surface of the sheath by a highly elastic cellular tissue ; the chord maintains its ropelike 
character along the basal third of the glans (lb. F), then subsides, expanding laterally, 
and is finally lost upon the firm capsule of the glans. There is no { os penis ' 

The nerves of the dorsum penis, the arteries, and trunks of two large plexuses of 
veins, pass beneath the bridge formed by the confluence of the tendinous and muscular 
parts of the 'levatores penis* and between the two suspensory ligaments of the penis. 
These ligaments are an inch in breadth, and one-third of an inch in thickness at their 
origin from the ischio-pubic arch a little in advance of the ligamentous attachments of 
the crura corporis caveniosi. 

The total length of the undistended penis is three feet nine inches ; the circumference 
of the prepuce is one foot five inches, 

The external and constantly exposed firmer tegumentary part of the prepuce has been 
already described, and is figured in PL IX. fig. 4. 

The substance of the large reflected preputial fold of softer integument (PL IX. fig. 5, 
and PL XVIL fig. i, pr) is from half an inch to two-thirds of an inch in thickness, and 
consists of a moderately compact cellular corium, with a delicate epiderm, minutely rugose 
in the transverse direction, and perforate or punctate with the pores of the mucous 
follicles which are very regularly dispersed at intervals of about a quarter of an inch* 

The glans penis (lb, ib. gl) is a long and slender subeompressed cone with a truncate 
apex; it measured in its flaccid undistended state, one foot in length: the prepuce is 
reflected upon its base at the same transverse or circular line, and there is no free mi m. 
The apex (PL IX. figs, 5 & 6, and PL XVIL figs. 1, a & 3, a) is not simple, but resembles 




a mushroom on a thick peduncle (fig. 3, p) projecting from an excavation (lb. e, e) at 
the end of the gtans with a thin wall or border, like a second prepuce ; bat this is of 
the same structure as the rest of the firm surface of the glans 

On each side of the base of the glans, and rather towards its under part, there is a 
longitudinal thick oblong ridge or lobe (PI. IX. figs. 5 & 6, and PI, XVII, fig, 1, r, r) t 
three inches and a half in length, and eight lines in basal thickness ; the thick rounded 
free border of each lobe inclines downwards. 

Mucous follicles similar to those oo the under surface of the prepuce extend from the 
attachment of that fold along one half of the interspace of the lateral lobes. The depth 
of the preputial fold is least on the dorsal side of the glans. On each side of the base 
of the glans, near the dorsum, the follicles extend along a space of two inches from the 
root of the prepuce, but do not occur on the middle line. A narrow ridge commences 
in the median space of the ' dorsum glandis s J which increases in height as it advances 
forwards, and then subsides two inches from the border of the terminal or apical fu*sa. 
The projecting border of this fossa (PI, XVII. figs. 2 & 3, e, e) describes a compressed oval, 
and is attached to the pedunculated appendage (lb. p, a) by a process, like a fraenum 
(lb, fig. 3,/,/), continued upon the middle line of both the upper and under surfaces of 
the thick peduncle (/&. p): the fossa between this peduncle and the free external border 
is not less than two inches in depth on each side ; the upper or dorsal border of the fold 
is three times the breadth of the under one. The stem p of the terminal expanded discoid 
appendage a is subcompressed with an oval section, one inch in long diameter, where it 
supports the terminal disc two-thirds of an inch across. The disc a is ovate, one inch 
eight lines long by one inch across its broader inferior part, where it extends farthest 
from the supporting stem. The urethra (lb. u) is perforated in the middle line of the 
terminal disc between its middle and upper third. 

The lateral lobes (lb. tig. 1, r, r) consist chiefly of erectile tissue ; and all the parts 
of this singularly complex glans are much altered in size, and somewhat also in shape, 
during erection (see PI. IX. figs, -I, 5 & 6). 

Female Organs, — The ovaria are included within a large peritoneal sac (PL XV [I. 
o, p) communicating with the general abdominal cavity by an opening which is 
three inches wide. They are compact, oblong, flattened bodies, with a smooth surface, 
as might be expected from the immature age of the animal. The left ovarium measured 
three inches and a half in length, by two inches and a half in breadth : the right was 
somewhat smaller, The external capsule of the ovarium is stoat and unyielding, and 
the serous covering has the appearance of being strengthened by tendinous lines, one 
of which runs in a curved direction across the anterior part of the ovary, having other 
shorter lines diverging from it. The stroma ovarii is also dense. Three ovisacs or 
Graafian vesicles were dissected out of each ovarium : of these one was an inch in 
diameter, with very dense thick dark-coloured parietes; the rest had a diameter of 
two-thirds of an inch, with thinner coats. Their contents were examined with great 



care under the microscope, but the granular layer was evidently broken up by decom- 
position, and the ovulum was invisible. The animal had been dead a fortnight. 

The Fallopian tubes or oviducts commence by wide orifices having a richly fimbriated 
margin (PI. XVII. /,/) ; their diameter at the expanded end equals two-thirds of an inch, 
but gradually diminishes in size as the tube passes in a slightly tortuous course along 
the parietes of the ovarian capsuie towards the uterus ; just before they enter the cornu 
uteri their diameter does not exceed one-third of a line ■ they terminate in the extremity 
of the cornu upon a valvular protuberance about the size of a pea, which is divided 
into four or five processes (lb, p). The inner surface of the oviduct is augmented by 
short irregular longitudinal folds or processes of the lining membrane. Each oviduct 
is fourteen inches in length. 

The coniua uteri (16. u\ u) are each seventeen inches in length, and uniformly 
about an inch in diameter; their area is occupied by close-set longitudinal folds of the 
lining membrane, about a quarter of an inch in breadth, and having a wavy irregular free 
margin. There is no appearance of processes for the attachment of cotyledons. Where 
the cornua join the body of the uterus, the crenation or scalloping of the longitudinal 
folds becomes shorter and deeper. The length of the common uterus (lb, cu) is 
about one and a half inch. The surface of its lining membrane is smooth, and pre- 
sented, when first exposed t a leaden hue. The place of the * os tineas' is occupied by 
a complex and remarkable structure, which will be described as it was traced from the 
vagina towards the uterus. 

A large transverse semilunar fold (J6, /,/} projects from the upper and lateral 
parts of the vagina ; the upper and broadest part of the fold is one inch ; it gradually 
diminishes as it descends on each side, and the cresses are lost about four inches from 
the vaginal orifice, and about an inch and a half from the middle line of the lower sur- 
face* About an inch above this fold, or nearer the uterus, a second and smaller fold is 
formed, which also descends from the upper and lateral walls of the vagina, but passes 
across in an oblique direction. Then follow in quick succession a series of shorter, 
but equally broad semilunar folds (Ib*f\f), which become alternate in their relative 
position as they approach the uterus, so as to cause the vagina to assume a spiral course 
not very unlike the disposition of the intestine in the Shark. As these valvular folds 
also assume a thicker, softer, and more vascular texture as they approach the uterus, 
it is by no means easy to determine where the vagina ends or the uterus begins. 
Measuring from the thickest fold which most resembles an os tincce, the common uterine 
cavity does not exceed two inches in length ■ and in this short extent, compared with 
the cornua, the Rhinoceros resembles the Elephant, 

The form, size, and relative positions of the vulval and preputial orifices have been 
already described. The length of the common or urogenital canal [lb, v, ug) was 
four inches ; its diameter about three inches. On each side, and about one inch and a 
half from the external outlet* were situated the apertures of the Malpighian or mucous 



canals (Ih. m, m). The diameter of the orifice was about a line, but as the canals 
passed inwards in the parietes of the urogenital passage, they widened to the diameter 
of from two to three lines. At about three inches, distance from their outlet they 
branched off into two or three smaller divisions, from which a glairy mucus could be 
expressed, and these again subdivided and terminated in blind secerning caeca applied 
to the outside of the commencement of the vagina. The orifice by which the vagina (w) 
communicates with the uro-genital canal is small in proportion to the width of that 
canal, and its area is diminished by several short oblique longitudinal folds, whose free 
edges project into it: the whole of this contracted orifice may he regarded as a form of 
hymen, beyond which the vagina rapidly dilates in the wide canal represented at v t v> 

Part V. 

Nervous System* 

Of this part of the Anatomy of the Rhinoceros the opportunities for making obser- 
vations were limited to the structure of the Brain and the Eye. 

The brain of the full-grown male (PL XIX. XX, & XXI.) weighed, when deprived of 
its membranes, 1 lb. 14| oz, avoirdupois, its proportion to that of the entire body being 
as 1 to Lu'4, 

An upper view of the natural size is given in PL XIX. fig, 1, a side view in fig. 2, 
and a base view in PL XX. 

The cerebral hemispheres present a subdepressed semioval form, broader behind, and 
narrower in front than in the Horse, and presenting fewer and larger con volutions. 
Their disposition resembles that in the larger hoofed mammalia generally ; converging 
from behind forwards as far as the anterior third of the cerebrum, and thence diverging 
as they extend forwards, but in a minor degree than in the Horse or Ox, 

In the view of the base of the brain (PL XX.), the large external crura jp, and internal 
crura 5, of the rhinencephalon or olfactory ganglia, 1,1, are shown, together with the 
protuberance r, which lies between the two crura. The chiasraa of the optic nerves is 
shown at 2, 2 ; the infundibular base of the hypophysis cerebri at k\ the single mass 
representing the corpora candicantia at l } the crura cerebri at i, and the third pair of 
nerves at 3, 3. The obtuse apices of the ' protuberantise natiformes/ 0, are less broad 
than in the Ox, and more resemble the shape of the same parts in the Horse. 

The cerebellum shows the small lobes, called ' flocci, 1 at A; and the inferior con- 
volutions at g. The olivary tracts make a very slight prominence at d; the pyramidal 
bodies are better defined, but are crossed by some superficial transverse fibres near the 
pores/: and the ' corpora trapezoidea/ e, are defined- The inferior ' vermiform process ' 
oi the cerebellum is more regular and better defined than the superior one : it is shown 
at v, tig, 4, PL XXII. 




The relative longitudinal extent of the great commissure or ' corpus callosum ' is 
shown at *, s, in the view of the vertical section of the brain of the female Rhinoceros 
given in PL XXII. fig* I. A septum lucidum, q f of moderate extent, connects the under 
surface of the anterior half of the corpus callosum with the fornix: in the same section 
the optic thalami are seen at o; the 'plexus choroides' at p\ and the mass of the 
tjuadrigeminal bodies at fc* The arrangement of the grey and white matter in the lobes 
and lobules of the cerebellum, forming the * arbor vitas/ a, is also shown : this is less 
complex than in the Horse. 

The lateral ventricle is laid open by the removal of its outer wall to show the size 
and shape of the great hippocampus at n, fig. 2, PL XXII. ; and in the same figure are 
shown the * plexus choroides * />, passing through the { foramen Monro! aim in * m, beneath 
the crura of the fornix and the outer lamina of the septum lucidum, q* The left lateral 
ventricle is laid open from above to show the proportions of the ( corpus striatum '/, 
with the hippocampus i t and the intervening part of the fornix, covering the optic tha- 
lamus h, together with the ' plexus choroides J p. In PL XXL the corpus callosum $, s, 
has been bisected and the hemispheres divaricated to show the forms and proportions 
of the bigeminal bodies ; of which the posterior pair 6 are broader but shorter than 
the anterior ones a. The pineal gland is shown at n; the optic thalamus at h\ the 
* plexus choroides ' at p ; and the ' corpus striatum ' at/. 

The commencing decomposition of the inner substance of the brain prevented the 
better definition, of some of the other parts of this organ. 

The common anastomotic trunk of the basilar or vertebral arteries, after traversing 
the median line of the pons, gives off a pair of arteries at right angles, which cross the 
crura cerebri between the pons and the third pair of nerves : a second pair of transverse 
branches is sent off" just anterior to the former, and receive the anastomosing longitu- 
dinal branches from the ento-carotids which complete the circle of Willis. From the 
ento-carotid parts of the circle, a branch is given off to the interspace between the 
middle and anterior lobes of the cerebrum, where it divides into three or four branches. 

The eyeballs are of small comparative size ; each measured in anteroposterior 
diameter one inch five lines, and in transverse diameter one inch three lines. Some 
dark-brown pigment lies under the conjunctiva for the extent of about a line from the 
circumference of the cornea : the same kind of pigment is also deposited upon the 
outside of the nictitating eyelid, and over a great part of the inner surface of the same 
part, covered of course by a reflection of the conjunctiva. The trunks of the vense 
vorticosa: perforate the sclerotica half-way between the entry of the optic nerve and 
the edge of the cornea. I injected one of these veins with mercury, which immediately 
returned by vessels perforating the sclerotica near the optic nerve. The disposition of 
the veme vorticosee, with the flocculent but somewhat firm connecting tissue of their 
radiating branches, presented that structure which most nearly resembled the figures 
given by Mr. Thomas of the parts he describes as " processes having a muscular 


r - 

appearance, with the fibres running forwards in a radiated direction 1 ." There are no 
fibres accompanying the radiated branches of the veins, showing the striated character 
of voluntary muscle under the microscope, Mr. Thomas found that e *the ciliary pro- 
cesses were affixed to the crystalline lens ; " but on removing the anterior part of the 
sclerotica, whilst the eye was suspended in spirit, both the vitreous humour and the 
lens rolled out ; and the capsule of the lens showed oo particular mark of the insertion 
or fixation of the ciliary processes ; their impressions, in remains of pigmental matter, 
were perceptible on the anterior part of the canal of Petit. The transverse diameter of 
the lens was six lines, the anteroposterior diameter four lines. Mr. Thomas also states 
that " the pigment was confined to the inside of the choroid 2 /' But in both Rhinoceroses 
dissected by me, I found on the outside of the chorion much loose cellular tissue, with 
dark pigment: this coloured flocculent tissue concealed at first the venae vorticose, 
even when injected. The sclerotica is one line thick at the back part of the eyeball ; 
and is thinnest near the middle of the ball, becoming thicker towards the cornea, which 
is two lines thick. The choroid adheres pretty strongly to the back part of the sclerotic, 
around the entry of the optic nerve, both by the entering vessels and by the tenacity of 
its outer flocculent coat, especially where the vessels penetrate the sclerotica. There 
is no tapetuni lncidum. The lower eyelid has a special depressor muscle. 


All the parts are of the natural size except when otherwise expressed. 


Fig. 1, Metacarpal gland* 

Fig, 2, Metatarsal gland, laid open. 

Fig. 3. Excretory orifice of the gland, 

Fig. 4, External prepuce during the ordinary retracted state of penis {one-sixth 
natural size). 

Fig. 5, Penis as protruded when this retromingent quadruped stales (one-sixth na- 
tural size). 

Fig. fi. Gtans penis, or the portion uncovered by the prepuce, when the organ is in 
a state of erection (one-sixth natural size). 

Fig, 7. Externa! parts of generation in the female (two-thirds natural size). 

Fig. 8. The two teats. 

Fig. 1. Right tonsil, epiglottis, arid backpart of larynx, 

1 Fijiloso^Wul Transactions, 1S0L p. 150. pi. 10- figs. 1, % 5. 

* lb, p. 150- 



Fig, 2. Inner surface of left side of the larynx laid open ; a part of the accessory fold 
of membrane t, and of the lower c vocal chord ' v t of the right side are pre- 
served and turned forwards : f t accessory fibre-cartilage ; u f upper ' vocal 
chord ' ; tk, thyroid cartilage ; cr , cricoid cartilage ; tr, first tracheal ring ■ 
e, epiglottis. 


Fig. 1. Outer surface of the stomach, with the serous tunic partially reflected to show 
the two decussating muscular layers (one-eighth natural size). 

Fig, 2. Inner surface of the stomach (one-eighth natural size). 

Fig. 3. Section of the coats of the stomach at the junction of the epithelial with the 
gastro-mucous linings. 

Fig, 4, A portion of the inner surface of the stomach, showing the follicular structure 
at the termination of the cardiac epithelium. 


Fig. 1. Portion of the inner surface of the beginning of the jejunum. 
Fig. 2. Portion of the inner surface at the end of the jejunum. 
Fig. 3. Portion of the inner surface near the end of the ileum. 


The caecum, colon, and beginning of the rectum (one-tenth natural size) 


Fig, 1. The inner surface of the duodenum, showing the terminal orifices of the 

biliary and pancreatic ducts. 
Fi g, 2. The inner surface of the fundus vesicae, showing the allantoic or urachal 

Fig, 3. Outside view of the right kidney (one-third natural size). 
Fig. 4, View of a portion of the pelvis of the kidney, with the beginning of the 



Fig. 1. Back view of the larynx, showing, f A, thyroid cartilage; c, cricoid cartilage; 
a a, arytenoid cartilages ; e, epiglottis ; d, erico-arytenoidei muscles, the 



left reflected; o t the right anjtenoideus, reflected; e, tkyro- cricoid? us ; c, 
articular tubercle, with a synovial surface for the joint with the base of the 
arytenoid cartilage. 

Fig. 2. Back view of the larynx, showing in addition to the foregoing, — ft, the com- 
missural cartilage of the arytcnoidei muscles o o ; g, the right thyro-aryte- 
noideus ; m, the superior labia of the glottis ; n, the upper * chorda vocalis 1 ; 
v t the lower 'chorda vocalis' bounding the entry of the laryngeal sac ; /, the 
fibro-cartilage giving attachment to the epiglottideal fold of membrane I, 
which bounds the upper part oi the suprachordal sacculus* 


The urinary bladder, vasa deferentia, vesicular, prostatic, and Cowperian glands (one^ 
fifth natural size)* The letters are explained in the text. 


Fig, 1. Muscles of the penis: c, crura penis; pr, reflected prepuce; gl gl, glans 
penis ; r f lateral lobes of glans ; a t terminal appendage of glans ; I l 7 car- 
neous mass of levatores penis ; l f beginning, and l u insertion of the common 
tendon of the levatores, which runs along the * dorsum penis ' ; 1 1, retractores 

Fig, 2. Front view of terminal fossa, and appendage of the glans penis t u, the orifice 
of the urethra. 

Fig. 3, Oblique view of the same parts, showing, ee, the borders of the fossa ; p, the 
peduncle of a, the discoid appendage ; //, its upper and lower freena ; u, 
urethral orifice, 

Fig t 4. Part of the * cervix vesicae' showing the common orifices, into which bristles 
are inserted, of the vasa deferentia and vesicular glands, upon the crucial 
verumontanum ; and the smaller excretory pores of the prostatic lobes. 


The female organs of generation (one-fifth natural size) : op, ovarian capsules ; /, 
fimbriae of oviduct; p, valvular papilla on which the oviduct terminates in 
the uterus ; u ! u t cornua uteri, the left laid open ; a probe is passed through 
the coats of the right cornu to show the place of confluence of the cornua 
with the £ corpus uteri ' ; f j\ valvular folds of ' cervix uteri J and r fundus 



vaginae ' ; (the complex glans penis has, probably, relation to this structure ;) 
v v 3 vagina; u, the urethra, beyond the hymeneal constriction dividing the 
vagina from the urogenital canal ; m m $ Malpighian canals, the left laid 
open — bristles are inserted into both ; vu f vulva and clitoris. 


Brain of the male Rhinoceros. 


Base of the brain of the male Rhinoceros. 


Dissection of the brain of the male Rhinoceros. 


Dissections of the brain of the female Rhinoceros. The letters are explained in the 

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